Int. J. Plant Sci. 180(3):220–231. 2019. q 2019 by The University of Chicago. All rights reserved. This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License (CC BY-NC 4.0), which permits non-commercial reuse of the work with at- tribution. For commercial use, contact [email protected]. 1058-5893/2019/18003-0005$15.00 DOI: 10.1086/701468 EOCENE FOSSIL LEGUME LEAVES REFERABLE TO THE EXTANT GENUS ARCOA (CAESALPINIOIDEAE, LEGUMINOSAE) Patrick S. Herendeen1,* and Fabiany Herrera* *Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, Illinois 60022, USA Editor: Michael T. Dunn Premise of research. Fossil leaves from the early Eocene Green River Formation of Wyoming and late Eo- cene Florissant Formation of Colorado have been studied and described here as two species in the monospe- cific extant genus Arcoa (Leguminosae, subfamily Caesalpinioideae). The single living species of Arcoa is en- demic to the Caribbean island of Hispaniola. The species from Florissant has been known since the late 1800s but has been incorrectly treated as several different legume genera. Methodology. The compression fossils were studied using standard methods of specimen preparation and microscopy. Fossils were compared with extant taxa using herbarium collections at the Field Museum and Smithsonian Institution. Pivotal results. The fossil bipinnate leaves exhibit an unusual morphological feature of the primary rachis, which terminates in a triad of pinnae, one terminal flanked by two lateral pinnae, all of which arise from the same point at the apex of the rachis. This feature, combined with other features that are diagnostic of the family Leguminosae or subgroups within it, allows the taxonomic affinities of the fossil leaves to be definitively deter- mined as representing the extant genus Arcoa, which is restricted to the Caribbean island of Hispaniola today. Conclusions. The fossil leaves described in this article demonstrate that the monospecific genus Arcoa was more diverse and had a much more widespread distribution in the past than it has today. Although the two fossil species are clearly referable to the same genus, differences between them in leaf size are consistent with differences in climate that are inferred for the more tropical Fossil Lake flora of the Early Eocene Green River Formation as compared with the warm temperate Late Eocene Florissant flora. Keywords: Arcoa, Caesalpinioideae, Caribbean, fossil, Hispaniola, Leguminosae. Introduction Wang et al. 2006, 2007, 2014; Bruneau et al. 2008; Wing et al. 2009; Collinson et al. 2012; Pan et al. 2010, 2012; Jia and Man- The legume family (Leguminosae, Fabaceae) is the third- chester 2014).Whilemanyfossillegumeleaves,flowers,andfruits largest family of flowering plants (ca. 768 genera, 19,600 spe- have been identified to genus, many more remain unidentified cies) and is ecologically very important in almost all terrestrial (Herendeen et al. 1992). In most cases, the unidentified dispersed habitats (Lewis et al. 2005; G. P. Lewis, personal communica- fossil leaves or fruits lack distinctive or diagnostic features (Lewis tion). The family has an abundant fossil record, beginning in the et al 2005; LPWG 2017) that would allow them to be referred to late Paleocene and diversifying rapidly through the Eocene (He- one or more particular groups within the family; they are clearly rendeen et al. 1992). By the late Eocene the fossil record of the legumes but not distinctive in any way that would aid identifica- family is diverse, with numerous caesalpinioid, mimosoid, and tion. In other cases, the fossils may exhibit distinctive features papilionoid taxa documented from many fossil localities around (e.g., winged fruits, bipinnate leaves terminating in a triad of pin- the world. The fossil record of the family was reviewed in the nae, etc.), but in such a diverse family with ca. 768 genera it can book Advances in Legume Systematics, Part 4 (Herendeen and takeconsiderabletimetodeterminewhichcladeswithinthefamily Dilcher 1992), and subsequently expanded on by numerous ad- share these characters. The latter situation applies to the fossils ditional contributions (e.g., Burnham 1995; Herendeen and described in this article. Jacobs 2000; Calvillo-Canadell and Cevallos-Ferriz 2002, 2005; In this article, we describe two species of bipinnate legume leaves from the Eocene of Colorado (early Eocene Green River fl fl 1 Author for correspondence; email: pherendeen@chicagobotanic Fossil Lake ora, late Eocene Florissant ora) and discuss their .org. systematic relationships within the family. The species from ORCIDs: Herendeen, https://orcid.org/0000-0003-2657-8671; Herrera, Florissant was previously described, first as a species tentatively https://orcid.org/0000-0002-2412-672X. assigned to the extant genus Caesalpinia by Lesquereux (1874) Manuscript received July 2018; revised manuscript received October 2019; elec- and subsequently as a speciesinthe fossil genusMimositesBower- tronically published February 5, 2019. bank (Lesquereux 1878) and later transferred by MacGinitie 220 HERENDEEN & HERRERA—EOCENE FOSSIL LEGUME LEAVES REFERABLE TO ARCOA 221 (1953) to the extant genus Prosopis L. (Mimosoideae). The fossil Collections of the US National Museum of Natural History, leaves from Green River and Florissant are unusual in that they Smithsonian Institution. The specimens have been cited in Les- are bipinnate and terminate with a triad of pinnae, a central ter- quereux (1874, 1878) and Knowlton (1898) and described in minal pinna, and a pair of lateral pinnae, all attached at the apex some detail in MacGinitie (1953). of the primary rachis. This is in contrast to most taxa with bipin- Fossil specimens were studied using light microscopy and nate leaves, in which the leaf terminates with a pair of pinnae. photographed using a Canon Rebel camerawith100-mm macro- Although the leaves and leaflets have a stereotypical mimosoid lens attached to a Stackshot system, and digital images were appearance, and many extant mimosoid genera have bipinnate merged using Helicon Focus software. Herbarium specimens leaves, no extant mimosoid genus is characterized by the pres- from the Field Museum (F) and National Museum of Natural ence of bipinnate leaves that terminate with a triad of pinnae. History, Smithsonian Institution (US), were used for compari- Leaveswiththisarchitecturedooccurintwodistinctclades within son with the fossils. the Caesalpinioideae. Although it is straightforward to make a list of all genera and species known to have bipinnate leaves, Results the distinctive feature that characterizes the fossils described here is not so easily investigated because it is not always mentioned in The two species treated here share a number of morpholog- genus or species descriptions. Thus, specimens often need to be ical features in common, including bipinnate leaf structure, op- examined in herbarium collections to document this feature. posite pinnae and leaflets, essentially sessile leaflets, and a fleshy The fossil plants are from two Eocene age localities, the Fossil transversely wrinkled pulvinus at the petiole base, at the bases Lake flora from the Fossil Butte Member of the Green River For- of pinnae, and on the leaflet bases. The most distinctive and di- mation (lower Eocene of Wyoming) and the Florissant Forma- agnostic feature exhibited by both of these species is that the tion (upper Eocene of Colorado). The fossils from the Green leaf terminates with three pinnae, of which two are lateral River Formation described here occur in limestone deposits and one is central or terminal. The number of pairs of pinnae from Fossil Lake in southwestern Wyoming. Fossil Lake was the and number of leaflets per pinna differ between the two species, smallest of three lakes that make up the Green River Formation. as do leaflet size and overall size of the leaves. As a consequence, Fossil Lake is the source of a diverse biota of fossil animals and we recognize these fossils as two species belonging to the same plants, which have been studied for many years. Grande (2013) genus. provides a history of Fossil Lake and descriptions of the animals and plants that have been documented from the rich fossil de- Systematics posits of this basin. The fossils from the Florissant Formation were collected in the later 1800s and early 1900s and described in Family—Leguminosae several publications, including Lesquereux (1874, 1878, 1883), Knowlton (1898), and MacGinitie (1953). The Florissant fossil Subfamily—Caesalpinioideae locality is now preserved at the Florissant Fossil Beds National Monument in Florissant, Colorado. Fossil plants, insects, and Genus—Arcoa Urb. vertebrates are preserved in lacustrine sediments of a relatively small lake that formed during the early stages of uplift of the Type—Arcoa gonavensis Urb. Rocky Mountains. Veatch and Meyer (2008) provides a history of paleontological research on the Florissant deposit. These fos- The genus Arcoa, with its single living species, is composed sil leaves are interesting because they demonstrate that Arcoa of small trees with two types of leaves, bipinnate leaves that ter- was more diverse and had a much more widespread distribution minate in a triad of pinnae, and once-pinnate leaves. Both leaf during the Eocene. Arcoa today includes a single living species types are often found on the same branch. Flowers are unisexual endemic to Hispaniola. The genus belongs to the Umtiza clade, (plants dioecious). Leaves and flowers are borne on short shoots.